Integrative taxonomy of the Lauraceae - feeding species of the genus Stephanitis (Hemiptera, Heteroptera, Tingidae) from Japan Author Souma, Jun https://orcid.org/0000-0002-2238-5015 Entomological Laboratory, Graduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, Fukuoka, Japan & Research Fellowship for Young Scientists (DC 1), Japan Society for the Promotion of Science, Tokyo, Japan kodokusignal@gmail.com text Deutsche Entomologische Zeitschrift 2022 2022-12-15 69 2 219 281 http://dx.doi.org/10.3897/dez.69.89864 journal article http://dx.doi.org/10.3897/dez.69.89864 1860-1324-2-219 BFE2BE4759E9450A807079B702A38625 9EE9E317E8195E3A94756A0DBBB1C35B Stephanitis (Norba) exigua Horvath , 1912 [Japanese name: Himetabu-gunbai] Figs 2C , 4C , 7C , 9C , 11C , 13C , 15C , 17C , 19C , 21C , 23C , 25C , 27C , 29C , 31C , 35 , 40H, I Stephanitis (Norba) exigua Horvath , 1912: 336. Syntype(s) (Fig. 34C ): Japan: Okinawa [= Ryukyu Islands, Okinawa-ken (an administrative area including a number of islands) or Okinawa Island (an island)] and Tateyama [= Honshu, Chiba-ken, Tateyama-shi, former Tateyama-machi in early 20th century (current Tateyama, Kamisanagura and Shimosanagura)]; ELHU and HNHM. Stephanitis (Norba) aperta Horvath , 1912: Azuma and Kinjo (1987 : 34) (distribution). Misidentification. References. Drake (1937 : 594) (distribution); Drake (1948 : 55) (checklist: Stephanitis ); Takeya (1951b : 13) (checklist: Japan); Drake and Maa (1953 : 100) (checklist: Stephanitis ); Takeya (1953 : 168) (distribution: part); Takeya (1963 : 38) (distribution: part); Drake and Ruhoff (1965 : 367) (catalog); Miyamoto (1964a : 274) (distribution: part); Miyamoto (1964b : 524) (distribution: part); Lee (1969 : 246) (male genitalia); Jing (1981 : 348) (monograph); Miyamoto and Yasunaga (1989 : 168) (checklist: Japan); Tomokuni (1994 : 843) (type material); Pericart and Golub (1996 : 58) (catalogue: Palaearctic; Yamada and Tomokuni (2012 : 204) (monograph: part); Yamada and Ishikawa (2016 : 433) (checklist: Japan); Nakatani (2021 : 78) (distribution). Material examined. Syntype ( 1 ♀ , ELHU) (Fig. 34C ), JAPAN : Honshu: "タテヤマ" [= Chiba-ken, Tateyama-shi, former Tateyama-machi in early 20th century (current Tateyama, Kamisanagura and Shimosanagura; approximate coordinates: 34°58'50.9"N , 139°51'27.1"E )], 11/VIII 1905 [= 11.viii.1905 ], "Matsumra" [sic; = collected by Shonen Matsumura and/or deposited in Matsumura's collection]. As pointed out in a previous study ( Tomokuni 1994 ), this single female syntype corresponds to Stephanitis (Stephanitis) pyrioides ( Scott , 1874) and does not match the original description of S. (Norba) exigua ( Horvath 1912 ). Therefore , if the remaining syntypes are present in ELHU and/or HNHM, a lectotype should be designated. Nevertheless , the former curator of HNHM does not know if the syntype of S. (N.) exigua exists in the collection ( D. Redei , pers. comm. 2021). Suspected syntypes ( 2 ♂♂ 1 ♀ 1 nymph, ELHU) (Fig. 35 ), JAPAN : Ryukyu Islands : " Okinawa " [= Okinawa-ken (a prefecture including a number of islands) or Okinawa Island (an island)], "6 29". These four individuals were labelled with inscriptions of " Stephanitis yaeyamae " (unpublished name) and "Matsum" (collected by Shonen Matsumura and/or deposited in Matsumura's collection). The species epithet of the unpublished name seems to refer to the Yaeyama Islands , the southern part of the Ryukyu Islands , but this morphological species is only distributed in the Daito Islands and the central part of the Ryukyu Islands . The morphological characteristics and locality data of three adult specimens match the original description of S. (N.) exigua ( Horvath 1912 ). However , the collector data of the four specimens are unclear and syntype (s) from " Okinawa " was (were) collected by "Kuroiwa" ( Horvath 1912 ). Therefore , these three adults could correspond to syntypes of S. (N.) exigua . In the present study, the author identified S. (N.) exigua based on three adult individuals. Non-types ( 106 ♂♂ 189 ♀♀ 7 nymphs), JAPAN : Ryukyu Islands (central part): Okinawa Island : 10.viii.1957 , leg. T. Takara ( 1 ♀ , NSMT); Kin , 14.vi.1958 , leg. T. Takara ( 2 ♀♀ , NSMT); Osato , 8.xi.1960 , leg. K. Yasumatsu ( 2 ♂♂ 1 ♀ , ELKU; 3 ♀♀ , KUM); Yona , 14.xi.1960 , leg. K. Yasumatsu ( 1 ♂ , KUM); as above but 19.xi.1963 , leg. H. Hasegawa ( 1 ♀ , KUM); as above but 24.iii.1964 , leg. Y. Miyatake ( 1 ♀ , KUM); as above but 23.xi.1985 , leg. M. Hayashi ( 2 ♂♂ 1 ♀ , TUA); as above but 28.vi.1984 , leg. M. Tomokuni ( 2 ♀♀ , NSMT); Shuri , 2.vi.1961 , leg. O. Nakoshi ( 1 ♂ , KUM); Nago , 22.x.1963 , leg. S. Ueno ( 1 ♂ , KUM); Tamagusuku , 17.xi.1963 , leg. H. Hasegawa ( 1 ♀ , KUM); Kudeken , 20.iii.1964 , leg. Y. Miyatake ( 2 ♂♂ 4 ♀♀ , KUM); Izumi , 22.iii.1964 , leg. T. Shirozu ( 9 ♂♂ 24 ♀♀ , KUM); Izumi-Gogayama , 22.iii.1964 , leg. S. Kimoto ( 1 ♂ , KUM); as above but leg. Y. Miyatake ( 3 ♂♂ 2 ♀♀ , KUM); Shoshi , 23.iii.1964 , leg. S. Kimoto ( 1 ♂ 1 ♀ , KUM); Nago , 23.iii.1964 , leg. Y. Miyatake ( 1 ♂ 2 ♀♀ , KUM); Hiji-Yonahadake , 25.iii.1964 , leg. T. Shirozu ( 2 ♀♀ , KUM); Hiji-gawa , 26.iii.1964 , leg. T. Shirozu ( 1 ♀ , KUM); Chinen , Sefa utaki, 17.ii.1973 , leg. H. Hasegawa ( 1 ♀ , NIAES); Yona , 21.ii.1973 , leg. H. Hasegawa ( 1 ♀ , NIAES); Kunigami-son , Mt. Yonahadake , 29.vi.1984 , leg. M. Tomokuni ( 3 ♀♀ , NSMT); Hanejiokawa , 14.xi.1985 , leg. M. Hayashi ( 1 ♂ 2 ♀♀ , TUA); Kunigami , Mt. Nishime , 19.x.1987 , leg. M. Tomokuni ( 1 ♀ , NSMT); Kunigami , Hama , 20.x.1987 , leg. M. Sakai ( 1 ♂ 3 ♀♀ , NSMT); Kunigami , Ooguni-rindo , 8 km from Yona , alt. 300 m , 21.x.1987 , leg. M. Tomokuni ( 3 ♂♂ 3 ♀♀ , NSMT); Kunigami , Hiji-Hiji Fall , 22.x.1987 , leg. M. Tomokuni ( 1 ♂ , NSMT); Kudeken , Seifa-utaki , 8.x.1988 , leg. M. Sakai ( 1 ♂ 1 ♀ , NSMT); Kunigamison , 10-11.x.1988 , leg. K. Konishi ( 1 ♀ , NIAES); Nago City , 12.x.1988 , leg. K. Konishi ( 1 ♀ , NIAES); Nakijin , Uebaru , 23.x.1990 , leg. M. Hayashi et al. ( 1 ♂ , NSMT); Afuso , 3.iv.1991 , leg. M. Hayashi ( 1 ♂ , TUA); Mt. Terukubi , 5.v.1991 , leg. M. Hayashi ( 1 ♀ , TUA); Mt. Yonahadake , 3.iv.1999 , leg. M. Hayashi ( 1 ♀ , TUA); Hedo , 16.ix.2002 , leg. M. Hayashi ( 1 ♀ , TUA); Kisebaru , 10.xii.2010 , leg. M. Hayashi ( 1 ♀ , TUA); Manzamo , 30.iii.2013 , leg. M. Hayashi ( 1 ♂ , TUA); Nago , Katsuyama , Mt. Katsuudake , 9.vi.2015 , leg. H. Yoshitake ( 1 ♀ , NIAES); Motobu , Namizato , Yaedake-sakura-no-mori-koen , 30.iii.2018 , leg. H. Yoshitake ( 4 ♂♂ 4 ♀♀ , NIAES); Kunigami-son , Uka-rindo , 10.xi.2018 , leg. H. Yoshitake ( 1 ♀ , NIAES); Naha-shi , Shuri-sueyoshi-cho , Sueyoshi-koen , 5.i.2019 , leg. H. Yoshitake ( 1 ♀ , NIAES); Yaese Park , 19.i.2019 , leg. H. Shigetoh ( 4 ♂♂ 7 ♀♀ 1 nymph, TUA); as above but leg. H. Yoshitake ( 2 ♂♂ 1 ♀ , NIAES); Nago-shi , Tanodake , 28.iv.2019 , leg. R. Ito ( 1 ♂ , TUA); Kunigami-gun , Kunigami-son , Sate , 6.v.2019 , leg. R. Ito ( 1 ♂ 1 ♀ , TUA); Nago , Genka Shisen For. Rd. , 3.vi.2019 , leg. T. Saeki ( 1 ♂ 1 ♀ , TUA); Uruma-shi , Mt. Ishikawadake , 29.xii.2019 , leg. H. Shigetoh ( 2 ♂♂ 1 ♀ , TUA); Naha-shi , Ohnoyama Park , 8.iii.2020 , leg. J. Souma ( 4 ♂♂ 5 ♀♀ 3 nymphs); Kunigami-son , Aha , 5.v.2019 , leg. H. Yoshitake ( 1 ♀ , NIAES); as above but 18.iv.2020 ( 1 ♂ 1 ♀ , NIAES); as above but 18.iv.2020 , leg. H. Shigetoh ( 3 ♀♀ , TUA); Toyomigusuku , 6.xi.2020 , leg. J. Souma ( 1 ♀ , TUA); Tabaru , 6.xi.2020 , leg. J. Souma ( 1 ♂ 9 ♀♀ , TUA); Midorigaoka Park , 10.xi.2020 , leg. J. Souma ( 5 ♂♂ 4 ♀♀ 1 nymph, TUA); Nanjo-shi , Chinen-joseki , 30.i.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Kitanakagusuku-son , Taguchi , 3.ii.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Nakagusuku , Noborimata , 3.ii.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Okinawa-shi , Yaeshima-koen , 4.ii.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Uruma-shi , Enobi , 4.ii.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Yomitan-son , Zakimi-joseki , 5.ii.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Yonabaru-cho , Untamamori , 20.ii.2021 , leg. H. Yoshitake ( 1 ♀ , NIAES); Nago-shi , Makiya , 1-2.vi.2021 , leg. T. Saeki ( 1 ♀ , TUA). Aka Island : 13.viii.1977 , leg. M. Kinjo ( 1 ♀ , NSMT); as above but 14.viii.1977 ( 1 ♀ , NSMT); as above but 14.viii.1977 , leg. S. Azuma ( 2 ♂♂ 1 ♀ , NSMT); Aka , 4.v.2021 , leg. R. Ito ( 1 ♂ 3 ♀♀ , TUA). Fukaji Island : 3.v.2021 , leg. R. Ito ( 6 ♂♂ 6 ♀♀ , TUA). Geruma Island : 10.viii.1977 , leg. S. Azuma ( 1 ♀ , NSMT); 2.viii.2019 , leg. H. Shigetoh ( 1 ♀ , TUA); 4.v.2021 , leg. R. Ito ( 4 ♂♂ 2 ♀♀ , TUA). Kume Island : Une , Tonnaha-enchi , 27.iii.2018 , leg. H. Yoshitake ( 1 ♀ , NIAES); Daruma-yama , 28.iv.2018 , leg. R. Ito ( 3 ♂♂ 4 ♀♀ , TUA); Shirase-Riv. , 29.iv.2018 , leg. R. Ito ( 1 ♀ , TUA); Yamashiro , 22-24.vii.2020 , leg. H. Yoshitake ( 1 ♀ , NIAES). Tokashiki Island : near Shuraku , 27.iv.2019 , leg. H. Shigetoh ( 1 ♂ , TUA); Tokashiki , 7.xi.2020 , leg. J. Souma ( 2 ♂♂ 9 ♀♀ , TUA); Aharen , 7.xi.2020 , leg. J. Souma ( 2 ♂♂ 2 ♀♀ , ELKU; 1 ♂ 3 ♀♀ , TUA); Mt. Kumichizi , 8.xi.2020 , leg. J. Souma ( 2 ♂♂ 4 ♀♀ 2 nymphs, TUA); Otani-path , 1.v.2018 , leg. R. Ito ( 1 ♂ 2 ♀♀ , TUA); Tokashiki , Otani road, 30.iv.2021 , leg. R. Ito ( 1 ♀ , TUA); as above but 1.v.2021 ( 4 ♂♂ 4 ♀♀ , TUA). Tsuken Island : 16.iii.2019 , leg. H. Shigetoh ( 1 ♀ , TUA). Yabuchi Island : 5.iii.2020 , leg. J. Souma ( 1 ♂ 1 ♀ , TUA). Yagaji Island : Gabu , 9.iii.2020 , leg. J. Souma ( 2 ♂♂ 2 ♀♀ , TUA); Sumuide , 17.iv.2020 , leg. H. Shigetoh ( 1 ♂ , TUA). Zamami Island : near Mt. Odake , 20.iii.2020 , leg. H. Shigetoh ( 1 ♂ 3 ♀♀ , TUA); Asa Evacuation Route , 21.iii.2020 , leg. H. Shigetoh ( 1 ♀ , TUA); Ama , near Mt. Bansho , 21.iii.2020 , leg. H. Shigetoh ( 1 ♂ 1 ♀ , TUA); Inazaki-Kaminohama , 21.iii.2020 , leg. H. Shigetoh ( 1 ♀ , TUA); Ama , 3.vi.2020 , leg. H. Shigetoh ( 1 ♀ , TUA); Asa , 2.v.2018 , leg. R. Ito ( 2 ♂♂ 2 ♀♀ , TUA). Daito Islands : Kitadaito Island : Nakano , Daitogu Shrine , 24.xi.2021 , leg. T. Saeki ( 2 ♀♀ , TUA). Minamidaito Island : Ikenosawa , 10.iii.2013 , leg. H. Yoshitake ( 1 ♂ 1 ♀ , NIAES); Zaisho , 6.vii.2014 , leg. H. Ogai ( 5 ♂♂ 3 ♀♀ , TUA); Daito Shrine , 9.ii.2018 , leg. R. Ito ( 3 ♂♂ 2 ♀♀ , TUA). Six specimens from Aka and Geruma islands collected in 1977 were considered to be recorded as " Stephanitis aperta " by a previous study ( Azuma and Kinjo 1987 ) . Diagnosis. Stephanitis (Norba) exigua is recognised amongst other species of Stephanitis by a combination of the following characters: head, pronotal disc, marking on hemelytra and ventral surface in various shades of brown (Figs 7C , 9C , 11C , 13C , 15C , 17C , 19C , 21C , 23C ); calli light brown; body in male 2.2 times (in female 2.1 times) as long as maximum width across hemelytra (Figs 2C , 4C ); rostrum not reaching metasternum; pronotum unicarinate (Fig. 25C ); hood pale, shorter than median carina of pronotum, as wide as vertex at widest part, not covering eye, as high as median carina of pronotum at highest part, with posterior margin not extending to middle of pronotal disc; median carina of pronotum with 2 rows of areolae at highest part; pronotal disc lustrous; paranotum less erect, narrowed posteriorly, with 2 rows of areolae at widest part, with anterolateral angle slightly protruding anteriad, with outer margin gently curved inwards at posterolateral angle, maximum height shorter than height of eye (Fig. 27C ); apices of hemelytra close to each other in rest; costal area with 3 rows of areolae at widest part; subcostal area with 2 rows of areolae at widest part; discoidal area with 3 rows of areolae at widest part; sutural area with 3-4 rows of areolae at widest part; hypocostal lamina with a single row of areolae throughout its length; R+M (radiomedial) vein carinate; pygophore elevated at centre of venter, with posterior margin slightly emarginate in middle part (Fig. 29C ); and paramere stout, weakly curved inwards at apex, with outer margin not sinuate in middle part, inner margin nearly straight in basal part (Fig. 31C ). Remarks. Amongst the Japanese species of Stephanitis , S. (Norba) exigua is similar to S. (N.) hiurai in general habitus, but the former is easily distinguished from the latter by the following characters: hood as wide as vertex at widest part (wider than vertex in S. (N.) hiurai ), not covering eye (incompletely covering in S. (N.) hiurai ) (Figs 7C, E, F , 9C, E, F , 25C, E ); paranotum with 2 rows of areolae at widest part (3 rows in S. (N.) hiurai ), with anterolateral angle slightly protruding anteriad (protruding in S. (N.) hiurai ); costal area of hemelytron with 3 rows of areolae at widest part (4 rows in S. (N.) hiurai ) (Figs 15C, E, F , 17C, E, F ); subcostal area in female with 2 rows of areolae at widest part (3 rows in S. (N.) hiurai ); and R+M (radiomedial) vein in female carinate (not carinate in S. (N.) hiurai ). Distribution. Japan (Ryukyu Islands (central part): Okinawa Island, Aka Island, Fukaji Island, Geruma Island, Kume Island, Tokashiki Island, Tsuken Island, Yabuchi Island, Yagaji Island, Zamami Island; Daito Islands: Kitadaito Island, Minamidaito Island) (Fig. 45 ); China ( Horvath 1912 ; Drake 1937 ; Takeya 1963 ; Miyamoto 1964a , 1964b ; Azuma and Kinjo 1987 ; Yamada and Tomokuni 2012 ; Yamada and Ishikawa 2016 ; Nakatani 2021 ; present study). The previous record from Honshu ( Horvath 1912 ) is a misidentification of Stephanitis (Stephanitis) pyrioides . Hundreds of specimens from Honshu possessing the unicarinate pronotum were examined, but all of them belong to S. (Norba) aperta , S. (N.) mendica or S. (S.) tabidula . Therefore, S. (N.) exigua is probably not distributed in Honshu. The previous records from the southern part of the Ryukyu Islands and northern Taiwan ( Takeya 1963 ; Miyamoto 1964a , 1964b , 1964c ; Azuma and Kinjo 1987 ; Hayashi 2002 ) correspond to S. (Norba) ishikawai sp. nov., described below. Hundreds of specimens from the southern part of the Ryukyu Islands possessing the unicarinate pronotum were examined, but all of them belong to S. (N.) ishikawai sp. nov. Therefore, S. (N.) exigua is probably not distributed in the southern part of the Ryukyu Islands. In Japan, S. (N.) exigua inhabits the laurilignosa in a subtropical climate of the central part of the Ryukyu Islands, which is in the Oriental Region. Host plants. Cinnamomum camphora , "Kusunoki" ( Lauraceae ) ( Takeya 1963 ; present study); C. yabunikkei H.Ohba, "Yabunikkei" (present study); Machilus thunbergii , "Tabunoki" ( Lauraceae ) (Fig. 43E ) ( Takeya 1963 ; Yamada and Tomokuni 2012 ; present study). Stephanitis (Norba) exigua feeds only on lauraceous trees and is oligophagous. This lace bug sometimes occurs on plantings of C. camphora and M. thunbergii in its distribution range (present study), suggesting that it can become a pest of both lauraceous trees. Biology. Stephanitis (Norba) exigua feeds on the abaxial surface of leaves of the three host plants in Japan (present study). In Japan, adults were collected in almost all seasons ( Miyamoto 1964a , 1964b ; present study); nymphs were collected in January, March and November (present study).